Introduction

Polycystine Radiolaria is a major group of marine planktonic protists. Their skeletons are composed of amorphous silica, allowing their preservation in the sediments. In Costa Rica, radiolarians are the oldest fossils present in the geological record. Neogene radiolarians from the sedimentary cover of the Cocos plate are the subject of the present study. Their taxonomy and biostratigraphy is based on continuously cored deep-sea successions. Radiolarian data have been correlated with that of other microfossils, such as calcareous nannoplankton and foraminifers, and compared with geomagnetic polarity, in order to constrain the ages of the deposits.

During the last forty years, several authors have contributed to the development of regional stratigraphic radiolarian zonations for the Equatorial Pacific area (^{Riedel, 1957}, 1959; ^{Nigrini, 1971}; ^{Moore, 1971}; Riedel and Sanfilippo 1971, 1978; ^{Goll, 1980}). Radiolarians have also been used for paleoenvironmental reconstructions, identifying tropical, subtropical, cool temperate, polar and upwelling assemblages (^{Moore, van Andel, Sancetta & Pisias, 1978}). The association of radiolarian assemblages with water masses (differing chiefly in temperature and nutrients) has been described from many areas, including, among others, the Pacific Ocean (^{Nigrini, 1970}; ^{Riedel, 1971a}; ^{Sachs, 1973}; ^{Renz, 1976}; MolinaCruz, 1977; ^{Boltovskoy and Jankilevich, 1985}; Boltovskoy and Riedel, 1987), the Indian Ocean (^{Johnson and Nigrini, 1980}, ¹⁹⁸²) and the Atlantic Ocean (^{Bjørklund, Cotesea, Swanberg & Schrader, 1998}; Boltovskoy, 1998, ¹⁹⁹⁹). Thus, radiolarian fossil remains can also provide valuable clues to the ecological settings (primarily temperature and productivity) of the overlying waters at the time when these populations dwelled in the water-column. Although this technique, as most others, has its shortcomings (^{Boltovskoy, 1994}; ^{Haslett, 2002}), it has proven its usefulness in a numbers of detailed surveys world-wide.

Since the end of the 19th century, several high-level taxonomic schemes have been proposed for the polycystines. ^{Haeckel (1887}) produced one of the earliest comprehensive systems of radiolarian classification, describing thousands of new polycystine species. Haeckel’s work is still a necessary reference guide, but it does not satisfactorily represent natural relationships, because groupings are only based on morphologic similarities of the skeleton only, and because the rigidity of these geometry-based diagnoses often ignores the ample intraspecific variability of these protists. Efforts to improve the classification schemes inherited from earlier workers have followed two different main approaches, which are cytological data and evolutionary studies. Several revisions that rely heavily on cytoplasmic features have been proposed (^{Hollande and Enjumet, 1960}; ^{Cachon and Cachon, 1972a}, ^b; ^{Petrushevskaya, Cachon & Cachon, 1976}; ^{Petrushevskaya, 1981}). Although these schemes are probably sounder in biological terms, their application to fossil and subfossil materials lacking the protoplasm is problematic, which is one of the reasons for their very limited acceptance among radiolarian workers. Analyses of evolutionary lineages in geological sequences were used as a basis to assess the taxonomic value of key skeletal traits; it was concluded that many of them (e.g. number of segments, number of supplementary concentric spheres, number of feet, number of rays and of equatorial spines in discoidal Spumellaria, presence and nature of thoracic wings) have little or no suprageneric value. In contrast, several others (e.g. cephalic structure, pore arrangement, shell terminations in Nassellaria), traditionally considered as of minor value, are conservative through time and reveal evolutionary lineages and, therefore, are relevant for higher-rank divisions (^{Riedel and Sanfilippo, 1986}). Based on skeletal features only, several authors proposed alternative classifications, either for the entire order or for selected polycystine groups (Riedel, 1967a in ^{Banner et al., 1967}, 1971b; Petrushevskaya, 1965, 1971; ^{Goll, 1968}, 1969; ^{Sanfilippo and Riedel, 1970}; ^{Dumitrica, 1989}; ^{De Wever, Dumitrica, Caulet, Nigrini & Caridroit, 2001}). The presently most widely accepted classification for extant and Cenozoic radiolarians is still that of Riedel (1967a) and ^{Riedel (1971b)}, which is adopted in this work. The clasification of Miocene polycystines is based on De Wever et al. (2001).

This study is based on open ocean drill sites and intends to provide the first detailed MioceneRecent radiolarian inventory of Pacific sediments off Costa Rica. As opposed to most previous works in the area, especially those oriented at paleoceanographic reconstructions (see references above), which used restricted subsets of the fauna (usually 20-40 morphotypes), it covers all the radiolarians found in the samples.

Geological setting

The western active margin off Costa Rica represents the subduction zone resulting from the interaction of the Cocos plate and the Caribbean plate. The drilled sites of the expedition CRISP A2 are located near the interaction between the two plates, with the objective to characterize the rocks and sediments of the plate’s surrounds (Fig. 1). Hole U1381C is located about 4.5 km seaward of the accretionary prism, offshore the Osa Peninsula, on the incoming Cocos plate (8°25.7027′N, 84°9.4805′W, 2064.6 m water depth). Hole U1414 A is located on the incoming Cocos plate, about 1 km seaward of the deformation front (8°30.2304′N, 84°13.5298′W, 2459 m water depth) (Fig. 1). The hole U1381C is 109 m long and is dominated by pelagic and hemipelagic sediments intercalated by numerous tephra layers and a basaltic breccia layers at the bottom. The detail of the units is described by ^{Harris et al. (2013}) and ^{Schindlbeck et al. (2015}, ²⁰¹⁶). The Hole U1414 A measures 471.6 m, including 375.25 m of sedimentary deposits and 96.35 m of oceanic basaltic basement. The biogenic fraction mainly consists of calcareous nannofossil-rich ooze with foraminifers, diatoms, sponge spicules and radiolarians.

Material and methods

Two cores of the Integrated Ocean Drilling Program (IODP) Expedition 344: Costa Rica Seismogenesis Project, were sampled. The two cores contain calcareous nannofossil-rich ooze with foraminifera, diatoms and radiolarians. Two hundred and ninety-five samples were prepared (69 from Hole U1381C and 226 from Hole U1414 A) using the following standard procedure methods for Neogene sediments (^{De Wever, Dumitrica, Caulet, Nigrini & Caridroit, 2001}): (1) Sediments (10-20 g) were placed in beakers with 100-150 mL of 10% hydrogen peroxide. (2) Samples were boiled for a few minutes to achieve oxidation of the organic matter and disaggregation of the clay fraction. (3) Samples were rinsed with water and sieved through a 60 µm mesh. (4) The wet samples were placed again in beakers adding of a few drops of 10% hydrochloric acid to eliminate foraminifers, coccolithophorids, and other calcareous remains. (5) The samples were rinsed and sieved again through a 60 µm mesh. (6) A drop of the wet residue (1 mL) was placed onto a labeled glass slide, dried for a few minutes on a hot plate, embedded in Norland 60 mounting medium, and covered with a coverslip. (7) The slides were exposed for some minutes to ultraviolet light until dry. (8) The mounted samples were analyzed under a transmitted light microscope.

The age model was established using radiolarian data (^{Sandoval, Boltovskoy, Baxter & Baumgartner, 2017}), preliminary nannofossil shipboard data (^{Harris et al. 2013}) and published tephra dating with 40Ar/39Ar (^{Schindlbeck et al., 2015}, 2016).

Images of radiolarians were taken with an Olympus XC10 camera. In many cases, several photographs of the same specimen were obtained and processed with the aid of the Image J program (Extended Depth of Field), which allows merging a stack of photos in different focal positions into a single composite image (^{Forster, van de Ville, Berent, Sage & Unser, 2004}).

Systematic paleontology

Class ACTINOPODA^{Calkins, 1909}

Subclass RADIOLARIA^{Müller, 1858} Superorder POLYCYSTINA^{Ehrenberg, 1838}, emend. ^{Riedel, 1967}a in ^{Banner et al., 1967} Order SPUMELLARIA Ehrenberg, 1875, emend. Riedel, 1971a

Family COLLOSPHAERIDAE^{Müller, 1858}

Genus Acrosphaera^{Haeckel, 1881}

Acrosphaera sp. A (Fig. 2A)

Remarks: Regular sphere, covered with numerous, straight, radial spines, irregularly scattere over the whole surface. It presents some protuberances and depressions. Dimensions: ~100 μm in diameter, spines 20-40 µm.

Acrosphaera sp. B (Fig. 2B)

Remarks: Sub-circular sphere covered with numerous irregular pores. The size and shape of the spines is irregular. Dimensions: ~150-170 μm in diameter, spines 20-50 µm, pores size 10-50 µm.

Acrosphaera murrayana (Haeckel) (Fig. 2C) Choenicosphaera murrayana n. sp. Haeckel, 1887, p. 102, pl. 8, fig. 4.

Polysolenia murrayana (Haeckel) - ^{Nigrini, 1968}, p. 52, pl. 1, figs. 1a-b - Nigrini and Moore, 1979, pl. 2, figs. 4a-b.

Acrosphaera murrayana (Haeckel) -

^{Takahashi, 1991}, pl. 1, figs. 6-10.

Genus Siphonosphaera ^{Müller, 1858}

Siphonosphaera martensi Brandt (Figs. 2D and 2E)

Siphonosphaera martensi n.sp. ^{Brandt, 1905}, p. 339, pl. 9, figs. 9-12 - ^{Boltovskoy and Riedel, 1980}, p. 104, pl. 1, fig. 8 - ^{Takahashi, 1991}, p. 59, pl. 4, figs. 4-5, 7-8.

Remarks: Circular sphere covered with numerous irregular polygonal pores. Surface is gently sharped. Dimensions: ~100-130 μm in diameter, pores size 10-25 µm.

Siphonosphaera polysiphonia Haeckel (Figs. 2F and 2G)

Siphonosphaera polysiphonia n.sp. ^{Haeckel, 1887}, p. 106 - ^{Nigrini 1967}, p. 18-19, pl. 1, figs. 4a-b.

Genus Collosphera ^{Müller, 1855}Collosphaera macropora Popofsky (Figs. 2H, 2I, 2J and 2K)

Collosphaera macropora n.sp. ^{Popofsky, 1914}, p. 247, pl. 14, fig. 2 a-c - ^{Boltovskoy and Riedel, 1980}, p. 103, pl. 1, fig. 6 - ^{Takahashi, 1991}, p. 56, pl. 2, figs. 13-18.

Collosphaera sp.A (Fig. 2L)

Remarks: Spherical to sub-spherical sphere surrounded by numerous sub-circular pores, irregular arrangement of pores. In general the pores are small, it varies from ~20 to 5 µm. Few pointed spines. Dimension: ~150 µm in diameter.

Collosphaera elliptica Chen and Tan (Fig. 2M)

Collosphaera elliptica n.sp. ^{Chen and Tan, 1989}, p. 1, pl. 1, figs. 1-2 - Tan and Chen, 1999, p. 132, figs. 5-27.

Collosphaera tuberosa Haeckel (Fig. 2N)

Collosphaera tuberosa n.sp. ^{Haeckel, 1887}, p. 97 - ^{Molina-Cruz, 1977}, pl. 2, fig. 6 - ^{Boltovskoy and Riedel, 1980}, p. 104, pl. 1, fig. 7 - ^{Takahashi and Honjo, 1981}, p. 144, pl. 1, fig. 2.

Genus Otosphaera ^{Haeckel, 1887}

Otosphaera polymorpha Haeckel (Figs. 2O, 2P and 2Q)

Otosphaera polymorpha^{Haeckel, 1887}, p. 116, pl. 7, fig. 6 - ^{Nigrini and Moore, 1979}, p. S9, pl. 1, fig. 5.

Otosphaera aff. auriculata^{Haeckel, 1887}

(Figs. 2R and 2S) aff. ^{Haeckel, 1887}, p.116, pl.7, fig.5

Remarks: The shell bears three short tubes with perforated walls whose distal end is circular. Pores are very small, irregular shape and arrangement. Dimensions: ~100 µm, pores 1-5 µm.

Genus Trisolenia ^{Haeckel, 1881}

Trisolenia zanguebarica Ehrenberg (Fig. 2T) Trisolenia zanguebarica n.sp. Ehrenberg, 1872a, p. 321.

Family ACTINOMMIDAE^{Haeckel, 1862}, emend. ^{Riedel, 1967}a in ^{Banner et al., 1967}

Genus Actinomma ^{Haeckel, 1862}Actinomma langii (^{Dreyer) (Figs. 3A and 3B) Sphaeropyle langii Dreyer, 1889}, p. 89, fig. 54 - ^{Foreman, 1975}, p. 618, pl. 9, figs. 30-31 - ^{Suzuki, 2006}, p. 861, fig. 13.

Genus Cladococcus ^{Haeckel, 1860}Cladococcus cervicornis Haeckel (Fig. 3C) Cladococcus cervicornis Haeckel, 1860, p. 801 - ^{Boltovskoy and Riedel, 1980}, p. 108, pl. 2, fig. 5.

Genus Diartus^{Sanfilippo and Riedel, 1980}Diartus hughesi (Campbell and Clark) (Fig. 3D)

Ommatocampe (Ommatocampula) hughesi n.sp. ^{Campbell and Clark, 1944}, p. 23, pl. 3, fig. 12.

Diartus hughesi (Campbell and Clark) - ^{Sanfilippo and Riedel, 1980}, p. 1010 - ^{Nigrini and Sanfilippo, 2001}, p. 221.

Diartus petterssoni Riedel and Sanfilippo (Figs. 3E and 3F)

Cannartus (?) petterssoni Riedel and

^{Sanfilippo, 1970}, p. 520, pl. 14, fig. 3.

Diartus petterssoni (Riedel and ^{Sanfilippo) - Sanfilippo and Riedel, 1980}, p. 1010 - ^{Nigrini and Sanfilippo, 2001}, p. 223.

Genus Didymocyrtis ^{Haeckel, 1862}

Didymocyrtis penultima (Riedel) (Fig. 3G)

Panarium penultimum^{Riedel, 1957}, p. 76, pl. 1, fig. 1.

Didymocyrtis penultima (Riedel) - ^{Sanfilippo and Riedel, 1980}, p. 1010 - ^{Nigrini and Sanfilippo, 2001}, p. 241.

Didymocyrtis antepenultima (Riedel and Sanfilippo) (Fig. 3H)

Ommatartus antepenultimus n.sp. ^{Riedel and Sanfilippo, 1970}, p. 521, pl. 14, fig. 14.

Didymocyrtis antepenultima (Riedel and ^{Sanfilippo) - Sanfilippo and Riedel, 1980}, p. 1010 - ^{Nigrini and Sanfilippo, 2001}, p. 233.

Didymocyrtis laticonus (Riedel) (Fig. 3I) Cannartus laticonus n.sp. Riedel, 1959, p. 291, pl. 1, fig. 5.

Didymocyrtis laticonus (Riedel) - ^{Sanfilippo and Riedel, 1980}, p. 1010.

Didymocyrtis antepenultima - ^{Nigrini and Sanfilippo, 2001}, p. 237.

Didymocyrtis mammifera (Haeckel) (Fig. 3J) Cannartidium mammiferum n.sp. Haeckel,

1887, p. 375, pl. 39, fig. 16.

Cannartus mammiferus (Haeckel) - ^{Riedel, 1959}, p. 291, pl. 1, fig. 4.

Cannartus mammifer (Haeckel) - ^{Sanfilippo, Burckle, Martini & Riedel, 1973}, p. 216, pl. 1, fig. 7.

Didymocyrtis mammifera (Haeckel) -

^{Sanfilippo and Riedel, 1980}, p. 1010 - ^{Nigrini and Sanfilippo, 2001}, p. 239.

Didymocyrtis bassanii (Carnevale) (Fig. 3K)

Cannartidium bassanii n.f. ^{Carnevale, 1908},

p. 21, pl. 3, fig. 12.

Didymocyrtis bassanii (Carnevale) - ^{Kamikuri, Motoyama, Nishi & Iwai, 2009}, p. 729, fig. 9H.

Didymocyrtis tetrathalamus (Haeckel) (Fig. 3L)

Panartus tetrathalamus n.sp. ^{Haeckel, 1887},

p. 378, pl. 40, fig. 3.

Ommatartus tetrathalamus (Haeckel) - ^{Riedel and Sanfilippo, 1971}, p. 1588, pl. 1C, figs. 5-7.

Didymocyrtis tetrathalamus (Haeckel) - ^{Sanfilippo and Riedel, 1980}, p. 1010.

Didymocyrtis tetrathalamus tetrathalamus (Haeckel) - ^{Nigrini and Sanfilippo, 2001}, p. 246.

Genus Saturnalis ^{Haeckel, 1881}Saturnalis circularis Haeckel (Fig. 3M) Saturnalis circularis n.sp. Haeckel, 1887, p.

131 - ^{Nigrini, 1967}, p. 24, pl. 1, fig. 9.

Genus Druppactractus ^{Haeckel, 1887}

Druppotractus sp. (Fig. 3N)

Remarks: Ellipsoidal shell consisting of a thickwalled external shell and a thinner internal one.

Genus Hexacontium ^{Haeckel, 1881}Hexacontium armatum/hostile Cleve (Figs.

3O and 3P)

Hexacontium armatum n.sp. ^{Cleve, 1900}, p. 9, pl. 6, fig. 5 - ^{Boltovskoy and Riedel, 1980}, p.

112, pl. 10a-b.

Genus Stylosphaera ^{Ehrenberg, 1846}

Stylosphaera sp. (Fig. 3Q)

Remarks: Actinommid with two prominent three-bladed polar spines.

Genus Plegmosphaera ^{Haeckel, 1887}

Plegmosphaera spp. (Fig. 3R)

Remarks: A central sphere surrounded by an irregularly branched meshwork.

Family PHACODISCIDAE^{Haeckel, 1881}

Genus Heliodiscus ^{Haeckel, 1862}

Heliodiscus spp. (Fig. 3S)

Remarks: Arrangement and abundance of pores quite different from H. asteriscus. Spherical microsphere is laterally located.

Family COCCODISCIDAE^{Haeckel, 1862}

Genus Periphaena ^{Ehrenberg 1873}

Periphaena decora Ehrenberg (Fig. 3T)

Periphaena decora^{Ehrenberg, 1873}, p. 246; 1875, pl. 28, fig. 6 - ^{Riedel, 1957}, p. 258, pl. 62, fig. 1 - ^{Sanfilippo and Riedel, 1973}, p. 523, pl. 8, figs. 8-10; pl. 27, figs. 2-5.

Remarks: Spherical microsphere is apparently located in the centre of the medullary shell.

Family SPONGODISCIDAE^{Haeckel, 1862}

Genus Amphirhopalum ^{Haeckel, 1881}

Amphirhopalum ypsilon Haeckel (Fig. 4A)

Amphirhopalum ypsilon n.sp. ^{Haeckel, 1887}, p. 522 - ^{Nigrini and Moore, 1979}, p. S75-S77, pl.

10, figs. 1a-e.

Genus Dictyocoryne ^{Ehrenberg, 1860a}

Dictyocoryne truncatum (Ehrenberg) (Fig. 4B) Rhopalodictyum truncatum n.sp. Ehrenberg, 1861, p. 301.

Dictyocoryne truncatum (Ehrenberg) - ^{Nigrini and Moore, 1979}, p. S89, pl. 12, figs. 2a-b.

Dictyocoryne profunda Ehrenberg (^{Fig. 4}C) Dictyocoryne profunda n.sp. Ehrenberg, 1860a, p. 767 - ^{Nigrini and Moore, 1979}, p.

S87-S88, pl. 12, fig. 1.

Genus Euchitonia ^{Ehrenberg, 1860a}

Euchitonia furcata/elegans group

(Ehrenberg) (Figs. 4D and 4E)

Pteractis elegans n.sp. ^{Ehrenberg, 1872a}, p. 319; 1872b, p. 299, pl. 8, fig. 3.

Euchitonia elegans/furcata group?

(Ehrenberg) - ^{Boltovskoy, 1998}, fig. 15.70.

Genus Spongaster ^{Ehrenberg, 1860b}Spongaster tetras Ehrenberg (Fig. 4F) Spongaster tetras n.sp. Ehrenberg, 1860b, p. 833; 1872b, p. 299, pl. 4(3), fig. 8.

Spongaster tetras tetras Ehrenberg - Nigrini

and Moore, 1979, p. S93-S94, pl. 13, fig. 1 - ^{Boltovskoy, 1998}, fig. 15.67.

Spongaster sp. A aff. D. truncatum (Fig. 4G) aff. Rhopalodictyum truncatum Ehrenberg,

1861, p. 301

Remarks: Test with three spongy arms, having rounded corners

Genus Spongurus^{Haeckel, 1862}Spongocore puella Haeckel (Fig. 4H) Spongocore puella n.sp. Haeckel, 1887, p. 347, pl. 48, fig. 6 - ^{Nigrini and Moore, 1979}, p. S59, S61, S69, pl. 8, figs. 5a-c.

Genus Stylodictya ^{Ehrenberg, 1846}, 1847, emend. Kozlova in Petrushevskaya and

Kozlova, 1972

Stylodictya aculeata Jørgensen (Fig. 4I)

Stylodictya aculeata n.sp. ^{Jørgensen, 1905}, pl. 10, fig. 41a-c - ^{Boltovskoy, 1998}, fig. 15.62a.

Stylodictya validispina Jørgensen (Fig. 4J)

Stylodictya validispina n.sp, ^{Jørgensen, 1905}, p. 119, pl. 10, fig. 40 - ^{Nigrini and Moore, 1979}, pl. 13, fig. 5a-b.

Genus Spongodiscus ^{Ehrenberg, 1854}

Spongodiscus sp. (Figs. 4K and 4L)

Remarks: Test a spongy disc, thickened in the centre.

Genus Circodiscus Kozlova in^{Petrushevskaya and Kozlova, 1972}

Circodiscus spp. (Figs. 4M, 4N, 4O and 4P)

Remarks: circular to sub-circular disc, with several internal concentric rings. Sometimes presents a pylome.

Genus Spongurus ^{Haeckel, 1860}

Spongurus spp. (Figs. 4Q, 4R and 4S) Remarks: Oval-elongate spongy shell.

Genus Spongopyle ^{Dreyer, 1889}

Spongopyle osculosa Dreyer (Fig. 4T)

Spongopyle osculosa n.sp. ^{Dreyer, 1889}, p. 42, pl. 11, figs. 99-100 - ^{Nigrini and Moore 1979}, pl. 15, fig. 1.

Genus Spongotrochus ^{Haeckel, 1860}

Spongotrochus spp. (Figs. 5A, 5B and 5C)

Remarks: Spongy disc thickened in its central part, sometimes with spines.

Family LITHELIIDAE^{Haeckel, 1881}

Genus Larcopyle ^{Dreyer, 1889}

Larcopyle butschlii Dreyer (Fig. 5D)

Larcopyle bütschlii n. sp. ^{Dreyer, 1889}, p. 124, pl. 10,70 - ^{Molina-Cruz, 1977}, p. 335, pl. 3, figs. 10-11 - ^{Nigrini and Moore, 1979}, p. S131-S132, pl. 17, fig. 1b - ^{Boltovskoy, 1998}, figs. 5B-B’, 15.85.

Larcopyle nebulum Lazarus, Faust & Popova-

Goll (Fig. 5E and 5F)

Larcopyle nebulum n.sp. ^{Lazarus, Faust & Popova-Goll, 2005}, p. 111, pl. 5, figs. 1-13.

Genus Lithelius ^{Haeckel 1862}

Lithelius minor Jørgensen (Figs. 5G and 5H)

Lithelius minor n.sp. ^{Jørgensen, 1900}, p. 65, pl. 5, fig. 24 - ^{Boltovskoy and Jankilevich, 1985}, pl. 3, fig. 17.

Genus Larcospira ^{Haeckel, 1887}

Larcospira spp. (Figs. 5I and 5J)

Remarks: Shell consists of an oval spiral from a common origin.

Larcospira quandrangula Haeckel (Figs. 5K and 5L)

Larcospira quadrangula n.sp. ^{Haeckel, 1887}, p. 696, pl. 49, fig. 3 - ^{Nigrini and Moore, 1979}, p. S133, pl. 17, fig. 2 - ^{Takahashi and Honjo, 1981}, p. 150, pl. 6, fig. 2.

Family PYLONIIDAE^{Haeckel, 1881}, emend. ^{Dumitrica, 1989}

Genus Phorticium ^{Haeckel, 1881}Phorticium pylonium group Haeckel (Figs. 5M, 5N and 5O)

Phorticium pylonium n.sp. ^{Haeckel, 1887}, p.

709, pl. 49, fig. 10.

Genus Pylonium ^{Haeckel, 1881}

Pylonium sp. sensu Benson (Fig. 5P)

Pylonium sp. ^{Benson, 1966}, p. 250, pl. 16, fig. 2 - ^{Benson, 1983}, p. 507, pl. 6, fig. 5.

Genus Tetrapyle^{Müller, 1858}Tetrapyle octacantha group Müller (Fig. 5Q) Tetrapyle octacantha n.sp. Müller, 1858, p. 33, pl. 2, figs. 12-13; pl. 3, figs. 1-12 - ^{Nigrini and Moore, 1979}, p. S125, pl. 16, figs. 3a-b - ^{Takahashi and Honjo, 1981}, p. 150, pl. 6, figs. 5-6.

Spumell. gen and sp. indet. (Figs. 5R, 5S and 5T)

Order NASELLARIA^{Ehrenberg, 1875} Family SPYRIDAE Ehrenberg, 1846

Genus Amphispyris ^{Ehrenberg, 1872b}

Amphispyris reticulata (Ehrenberg) (^{Fig. 6}A) Dictyospyris reticulata n.sp. Ehrenberg, 1872b, p. 307, pl. 10, fig. 19.

Amphispyris reticulata (Ehrenberg) - ^{Boltovskoy, 1998}, fig. 15.95 - ^{Tan and Chen, 1999}, p. 279, fig. 5-198.

Genus Cantharospyris ^{Haeckel, 1887}

Cantharospyris platybursa Haeckel (Fig. 6B)

Cantharospyris platybursa n.sp. ^{Haeckel, 1887}, p. 1051, pl. 53, fig. 7 - ^{Takahashi and Honjo, 1981}, p. 152, pl. 7, fig. 32.

Genus Dendrospyris ^{Haeckel, 1881}

Dendrospyris bursa Sanfilippo and Riedel (Fig. 6C)

Dendrospyris bursa n. sp. ^{Sanfilippo and Riedel in Sanfilippo, Burckle, Martini & Riedel, 1973}, p. 217, pl. 2, figs. 9-13 - ^{Nigrini and Lombari, 1984}, p. N19, pl. 16, figs. 1a-f - ^{Kamikuri, Motoyama, Nishi & Iwai, 2009}, p. 736, fig. 14G.

Giraffospyris ^{Haeckel, 1887}, emend.

^{Goll, 1969}

Giraffospyris toxaria (Haeckel) (Fig. 6D)

Podocoronis (Dipocoronis) toxarium n.sp. ^{Haeckel, 1887}, p. 980, pl. 83, fig. 7.

Giraffospyris toxaria (Haeckel) - ^{Goll, 1969}, p. 335, pl. 56, figs. 1-2, 4, 7 - ^{Kamikuri, Motoyama, Nishi & Iwai, 2009}, p. 734, fig. 12P.

Genus Liriospyris ^{Haeckel, 1881}

Liriospyris parkerae Riedel and Sanfilippo (Fig. 6E)

Liriospyris parkerae n.sp. ^{Riedel and Sanfilippo, 1971}, p. 1590, pl. 2C, fig. 15; pl. 5, fig. 4 - ^{Kamikuri, Motoyama, Nishi & Iwai, 2009}, p. 734, fig. 12E.

Liriospyris spp. (Figs. 6F, 6G and 6H)

Remarks: Sagittal ring sub-circular; joined directly to front, apex, and back of lattice shell. Vertical spine very short, projecting from approximate midpoint of sagittal ring; no frontal or axial spines.

Genus Lophospyris ^{Haeckel, 1881}

Lophospyris pentagona pentagona

(Ehrenberg) Goll (Fig. 6I)

Ceratospyris pentagona n. sp. ^{Ehrenberg, 1872a}, p. 303, pl. 15, fig. 5.

Dorcadospyris pentagona (Ehrenberg) -

^{Goll, 1969}, p. 338-339, pl. 59, figs. 1-3.

Lophospyris pentagona pentagona (Ehrenberg) - ^{Nigrini and Moore, 1979}, p. N15-N16, pl. 19, fig. 5 - ^{Takahashi, 1991}, p. 102, pl. 28, figs. 9-14.

Genus Neophrospyris ^{Haeckel, 1887}Nephrospyris renilla Haeckel (Fig. 6J)

Nephrospyris renilla n.sp. ^{Haeckel, 1887}, p. 1101, pl. 90, figs. 9-10 - ^{Boltovskoy, 1998}, fig.

15.99.

Genus Androspyris ^{Haeckel, 1887}

Androspyris huxleyi (Haeckel) (Fig. 6K)

Lamprospyris huxleyi n.sp. ^{Haeckel, 1887}, pl. 89, fig. 14.

Androspyris huxleyi (Haeckel) - ^{Goll, 1980}, pl. 4, figs. 4-5.

Genus Zygocircus ^{Butschli 1882}, emend.^{Goll, 1979}

Zygocircus productus (Benson) (Fig. 6L)

Zygocircus productus n.sp. ^{Benson, 1966}, p. 288, pl. 19, figs. 14-15 - ^{Goll, 1979}, p. 381, pl. 2, figs. 4-9.

Genus Phormospyris ^{Goll, 1976}

Phormospyris sp. A (Fig. 6M)

Remarks: Bilocular cephalis, circular to elliptical pores. Sagittal ring asymmetrical, with a line of bigger pores compared to cephalis.

Phormospyris stabilis stabilis (Goll) (Fig. 6N)

Desmospyris anthocyrtoides^{Benson, 1966},

p. 332, pl. 23, figs. 6-8.

Dendrospyris stabilis^{Goll, 1968}, p. 1422, pl. 173, figs. 16-18, 20.

Phormospyris stabilis (^{Goll) stabilis Goll, 1976}, p. 390, pl. 1, figs. 1-13; pl. 2, figs. 7-14 - ^{Takahashi, 1991}, p. 104, pl. 30, figs. 2-5 - ^{Boltovskoy, 1998}, fig. 15.100.

Phormospyris stabilis capoi Goll (Fig. 6O)

Phormospyris stabilis capoi n.sp. ^{Goll, 1976}, p. 392, 394, pl. 5, figs. 1-2; pl. 6-7 - ^{Takahashi, 1991}, p. 104, pl. 29, figs. 15-18.

Phormospyris stabilis (Goll) scaphipes (Haeckel) (Fig. 6P)

Tristylospiris scaphipes n.sp. ^{Haeckel, 1887},

p. 1033, pl. 84, fig. 13.

Phormospyris stabilis scaphipes (Haeckel) - ^{Goll, 1976}, p. 394, pl. 8-9.

Tholospyris scaphipes (Haeckel) - MolinaCruz, 1977, p. 336, pl. 7, figs. 11-12.

Genus Tholospyris ^{Haeckel, 1881}

Tholospyris spp. (Figs. 6Q, 6R, 6S and 6T)

Remarks: Oval-shaped sagitally constricted skeleton with very heavy bars and circular to oval pores.

Family PLAGONIIDAE^{Haeckel, 1881}, emend. ^{Riedel, 1957}

Amphiplecta ^{Haeckel, 1887}

Amphiplecta acrostoma Haeckel (Fig. 7A)

Amphiplecta acrostoma n.sp. ^{Haeckel, 1887}, p. 1223, pl. 97, fig. 10 - ^{Petrushevskaya, 1971}, pl.

54, figs. 2-7.

Genus Cladoscenium ^{Haeckel, 1887}

Cladoscenium spp. (Fig. 7B, 7C and 7D)

Remarks: Cephalis large, sub-cylindrical with a rounded or dome-shaped top, with unequal, irregular pores of all shapes. Those specimens present a typical large horn above to the cephalis. The feet are three bladed.

Genus Clathrocanium ^{Ehrenberg, 1860a}, emend.^{Petrushevskaya, 1971}

Clathrocanium sp. (Fig. 7E)

Remarks: Cephalis dome-shaped to globular, with irregular pores. Thorax with three large gates each between adjacent feet; the distal margins of the gates defined by convex downward arched bars developed between adjacent feet. Those specimens present a typical large horn above to the cephalis.

Genus Lophophaena ^{Ehrenberg, 1847}

Lophophaena hispida (Ehrenberg) (Fig. 7F) Dictyocephalus hispidus n.sp. Ehrenberg 1872b, p. 202, 289, pl. 5, fig. 18.

Lophophaena hispida (Ehrenberg) -

^{Boltovskoy, 1998}, p. 66, fig. 15.109.

Lophophaena aff. hispida (Fig. 7G) aff. Dictyocephalus hispidus ^{Ehrenberg 1872b}, p. 202, 289; pl. 5, fig. 18

Remarks: Cephalis spherical, with small pores, thin spines. Thorax conical, its pores decreasing in size toward the base.

Lophophaena aff. capito (Fig. 7H) aff. Lophophaena capito^{Ehrenberg, 1873}, p. 242; 1875, pl.8, fig.6

Remarks: Cephalis spherical, with medium pores, thin spines and a central robust spine in the center of the cephalis.

Plagoniidae group (Figs. 7I, 7J, 7K and 7L)

Includes many generally similar forms with a lat-

ticed cephalis and with or without a rudimentary thorax. The systematics of these sometimes extremely abundant forms is confused, and they are generally ignored in most surveys. They include species cited under a variety of generic names, such as Amphiplecta, Arachnocorallium, Arachnocorys, Ceratocyrtis, Dimelissa, Lophophaenoma, Micromelissa,

Peromelissa, Psilomelissa, etc. (^{Boltovskoy, 1998}).

Pteroscenium spp. (Fig. 7M)

Shell campanulate with irregular circular pores. Horn slender pyramidal. Three divergent feet twice as long as the horn strongly curved, thorny, three-edged.

Genus Pteroscenium ^{Jørgensen, 1905}

Pteroscenium pinnatum Haeckel (Fig. 7N)

Pteroscenium pinnatum n.sp. ^{Haeckel, 1887}, pl. 53, figs. 14, 16 - ^{Takahashi, 1991}, p. 114, pl.

36, figs. 8-9.

Genus Tetraphormis (^{Haeckel), 1887}

Tetraphormis dodecaster (Haeckel) (Fig. 7O) Sethophormis dodecaster n.sp. Haeckel,

1887, p. 1248, pl. 56, fig. 12.

Tetraphormis dodecaster (Haeckel) -

^{Takahashi, 1991}, p. 108, pl. 32, fig. 7.

Clathrocorys aff. teuscheri (Fig. 7P) aff. Clathrocorys teuscheri^{Haeckel, 1887}, p. 1220, pl. 64, fig.10

Remarks: it consists of a three-bladed central spine, which bifurcates in two perpendicular spines tree times along the central spine.

Genus Helothus ^{Jørgensen, 1905}

Helothus histricosa Jørgensen (Figs. 7Q and 7R)

Helotholus histricosa^{Jørgensen, 1905}, pl. 16, figs. 86, 88 - ^{Boltovskoy and Riedel, 1987}, pl. 4, fig. 4 - Boltovskoy, 1998, 15.113 (a-c).

Helothus aff. histricosa (Fig. 7S) aff. Helotholus histricosa^{Jørgensen, 1905},

pl. 16, figs. 86, 88

Remarks: Test broad, conical, with generally a broadly rounded, partially hidden cephalis with two robust spines. The pores follow an arrangement compared to H. histricosa.

Sethophormis ^{Haeckel, 1881}

Sethophormis sp. A (Fig. 7T)

Remarks: Shell in the form of a Chinese peasant hat, thorax with a very delicate lace-like meshwork of irregular pores decreasing in size toward the periphery and many radial sinuous ribs. Shell diameter: >300 µm.

Family THEOPERIDAE^{Haeckel, 1881},

emend. ^{Riedel, 1967}a in ^{Banner et al., 1967}

Genus Artopilium ^{Popofsky, 1913}

Artopilium undulatum Popofsky (Fig. 8A)

Artopilium undulatum n.sp. ^{Popofsky, 1913},

p. 405-406, pl. 36, figs. 4-5.

Lophocorys undulata (Popofsky) - ^{Takahashi, 1991}, p. 120, pl. 40, figs. 9-10.

Genus Clathtrocyclas ^{Haeckel, 1881}

Clathrocyclas alcmenae Haeckel (Fig. 8B)

Clathrocyclas alcmenae n.sp. ^{Haeckel, 1887},

p.1388, pl. 59, fig. 6.

Clathtrocyclas monumentum (Haeckel) (Fig. 8C) Calocyclas monumentum n.sp. Haeckel,

1887, p. 138.5, pl. 73, fig. 9 - ^{Takahashi, 1991}, p.

112, pl. 34, figs. 9-11.

Genus Corocalyptra ^{Haeckel, 1887}

Corocalyptra kruegeri Popofsky (Figs. 8D,

8E and 8F)

Corocalyptra kruegeri n.sp. ^{Popofsky, 1908}, p. 289, pl. 35, fig. 8 - ^{Boltovskoy, 1998}, p. 72, fig. 15.136.

Corocalyptra sp. (Fig. 8G)

Remarks: Cephalis with an apical horn. Thorax large, campanulate, with regular, polygonal pores in transversal rows increasing in size distally. Abdomen restricted to a narrow brim with several rows of pores.

Corocalyptra cervus (Ehrenberg) (Fig. 8H)

Eucyrtidium cervus n.sp. ^{Ehrenberg 1872a},

p. 308.

Coracalyptra cervus (Ehrenberg) - ^{Takahashi, 1991}, p. 112, pl. 33, figs. 9-12 - ^{Boltovskoy, 1998}, p. 71, fig. 15.128.

Genus Cornutella ^{Ehrenberg, 1838}

Cornutella profunda Ehrenberg (Fig. 8I)

Cornutella clathrata (?) profunda n.sp. ^{Ehrenberg, 1854}, p. 241.

Cornutella profunda Ehrenberg - ^{Boltovskoy, 1998}, p. 71, fig. 15.127 - ^{Takahashi 1991}, 113, pl.35, fig.3-9.

Genus Cycladophora ^{Ehrenberg, 1846}

Cycladophora davisiana Ehrenberg (Figs. 8J and 8K)

Cycladophora (?) davisiana n.sp. ^{Ehrenberg, 1861}, p. 297.

Cycladophora davisiana Ehrenberg - MolinaCruz, 1977, pl. 7, fig. 19.

Theocalyptra davisiana (Ehrenberg) - ^{Nigrini and Moore, 1979}, pl. 24, figs. 2a-b.

Cycladophora davisiana Ehrenberg -

^{Boltovskoy, 1998}, fig. 15.131.

Genus Cyrtocapsella (^{Haeckel) 1887}

Cyrtocapsella cornuta (^{Haeckel) (Figs. 8L and 8M) Cyrtocapsa cornuta n.sp. Haeckel, 1887}, p. 1513, pl. 78, fig. 9.

Cyrtocapsella cornuta (Haeckel) - Sanfilippo and ^{Riedel, 1970}, p. 453, pl. 1, figs. 19-20 - ^{Kamikuri, Motoyama, Nishi & Iwai, 2009}, p. 731, fig. 10X.

Cyrtocapsella tetrapera (Haeckel) (Figs. 8N and 8O)

Cyrtocapsa tetrapera n.sp. ^{Haeckel, 1887}, pl. 78, fig. 5.

Cyrtocapsella tetrapera (Haeckel) -

Sanfilippo and ^{Riedel, 1970}, p. 453, pl. 1, figs. 16-18 - ^{Kamikuri, Motoyama, Nishi & Iwai, 2009}, p. 731, fig. 10Z.

Cyrtocapsella cylindroides (Principi) (^{Fig. 8}P) Sticocapsa cylindroides n.f. Principi, 1909, p. 20, pl. 1, fig. 66.

Lithocampe (Cyrtocapsella) cylindroides (Principi) - ^{Petrushevskaya, 1975}, p. 582, pl. 14, figs. 14-15.

Cyrtocapsella cylindroides (Principi) - ^{Nigrini and Lombari, 1984}, p. N103-104, pl. 23, fig. 2.

Cyrtocapsella japonica (Nakaseko) (Figs. 8Q, 8R and 8S)

Eusyringium japonicum n.sp ^{Nakaseko, 1963}, p. 193, pl. 4, figs. 1-3.

Cyrtocapsella japonica (Nakaseko) -

Sanfilippo and ^{Riedel, 1970}, p. 452, pl. 1, figs. 13-15 - ^{Kamikuri, Motoyama, Nishi & Iwai, 2009}, p. 731, fig. 10Y.

Cyrtocapsella sp. (Fig. 8T)

Remarks: Cephalis is spherical, cylindrical shape in general. Thorax is the same size of abdomen and post-abodominal chamber. The surface is rough. Pores are round and the approximately the same size in the entire specimen. ~150 µm of length.

Genus Dictyophimus Ehrenberg, 1842, emend.

^{Nigrini, 1968}

Dictyophimus infabricatus Nigrini (Figs. 9A and 9B)

Dictyophimus infabricatus n.sp. ^{Nigrini, 1968}, p. 56, pl. 1, fig. 6 - ^{Takahashi, 1991}, pl. 37, figs. 3-5.

Dictyophimus hirundo (Haeckel) (Fig. 9C) Pterocorys hirundo n.sp Haeckel, 1887, p. 1318, pl. 71, fig. 4.

Dictyophimus hirundo group (Haeckel) - ^{Nigrini and Moore, 1979}, pl. 22, figs. 2-4.

Dictyophimus spp. (Figs. 9D, 9E and 9F)

Remarks: Cephalis partly submerged into thorax, bears a large apical horn. Thorax pyramidal or conical, with three conspicuous legs.

Genus Eucecryphalus ^{Haeckel, 1860}

Eucecryphalus gegenbauri Haeckel (^{Fig. 9}G) Eucecryphalus gegenbauri n.sp. Haeckel, 1860, p. 836 - Haeckel, 1862, p. 308, pl. 5, figs. 12-15.

Theocalyptra gegenbauri (Haeckel) -

^{Boltovskoy and Riedel, 1980}, p. 126, pl. 5, fig. 18.

Eucecryphalus tricostatum (Haeckel) (Figs. 9H, 9I, 9J and 9K)

Theopilium tricostatum n.sp. ^{Haeckel, 1887}, p. 1322, pl. 70, fig. 6 - ^{Takahashi and Honjo, 1981}, p. 152, pl. 8, fig. 12 - ^{Boltovskoy, 1998}, p. 78, fig. 15.149.

Eucecryphalus spp. (Figs. 9L, 9M, 9N, 9O, 9P, 9Q, 9R, 9S and 9T)

Remarks: Shell with a small cephalis. Thorax widely open, with small, regularly arranged pores, circular proximally and becoming larger and more polygonal distally. Abdominal brim flat with pores. In some specimens the abdominal brim is not observe because is broken.

Genus Eucyrtidium ^{Ehrenberg, 1847}, emend.

^{Nigrini, 1967}

Eucyrtidium anomalum Haeckel (Fig. 10A)

Eucyrtidium anomalum n.sp. ^{Haeckel, 1862}, p. 323, pl. 7, figs. 11-13 - ^{Boltovskoy and Riedel, 1987}, p. 100, pl. 5, fig. 1 - Boltovskoy, 1998, p. 75, fig. 15.138.

Eucyrtidium calvertense Martin (Fig. 10B)

Eucyrtidium calvertense n.sp. ^{Martin, 1904}, p. 450-451, pl. CXXX, fig. 5 - ^{Chen, 1975}, p. 460, pl. 15, fig. 9.

Eucyrtidium cienkowskii Haeckel (Figs. 10C and 9D)

Eucyrtidium cienkowskii n.sp. ^{Haeckel, 1887}, pl. 80, fig. 9 - ^{Sanfilippo, Burckle, Martini & Riedel, 1973}, p. 221, pl. 5, figs. 7-11 - ^{Lazarus, 1990}, pl. 6, figs. 1-3.

Eucyrtidium hexagonatum Haeckel (Fig. 10E, 10F and 10G)

Eucyrtidium hexagonatum n.sp. ^{Haeckel, 1887}, p. 1489, pl. 80, fig. 11 - ^{Molina-Cruz, 1977}, p. 336, pl. 7, figs. 4-5 - ^{Takahashi, 1991}, p. 124, pl. 42, figs. 18-19.

Eucyrtidium hexastichus (Haeckel) (Fig. 10H) Lithostrobus hexastichus n.sp. Haeckel, 1887, p. 1470, pl. 80, fig. 15.

Eucyrtidium hexastichum (Haeckel) - ^{Boltovskoy and Riedel, 1980}, p. 124, pl. 5, fig. 10 - ^{Takahashi, 1991}, p. 125, pl. 42, fig. 22 - Boltovskoy, 1998, p. 75, fig. 15.139.

Eucyrtidium teuscheri Haeckel (Fig. 10I) Eucyrtidium teuscheri n.sp. Haeckel, 1887, p. 1491, pl. 77, fig. 5 - ^{Caulet, 1986}, pl. 5, figs. 1-8.

Eucyrtidium spp. (Figs. 10J, 10K, 10L, 10M, 10N, 10O, 10P, 10Q, 10R, 10S and 10T)

Remarks: Small spherical to sub-spherical cephalis with small apical horn. Thorax small, inflated, thick-walled. Abdomen and three-five post-abdominal segments thin-walled, with pores arranged in longitudinal rows.

Genus Lampromitra ^{Haeckel, 1887}

Lampromitra schultzei Haeckel (Fig. 11A)

Lampromitra schultzei n.sp. ^{Haeckel, 1887}, p. 1214, pl. 60, figs. 7-7a. - ^{Boltovskoy, 1998}, p.

63, pl. 15.115.

Genus Litharachnium ^{Haeckel, 1860}

Litharachnium tentorium Haeckel (Fig. 11B)

Litharachnium tentorium n.sp. ^{Haeckel, 1860}, p. 836; 1862, p. 281, pl. 4, figs. 7-10 - ^{Boltovskoy, 1998}, p. 75, fig. 15.148.

Genus Lipmanella ^{Loeblich and Tappan, 1961}

Lipmanella dictyoceras (Haeckel) (Figs. 11C and 11D)

Lithornithium dictyoceras^{Haeckel, 1860}, p. 840.

Lipmanella virchowii^{Boltovskoy and Riedel, 1987}, p. 100, pl. 5, fig. 6.

Lipmanella dictyoceras^{Takahashi, 1991}, p. 121, pl. 40, fig. 17 - ^{Boltovskoy, 1998}, fig. 15.134.

Lipmanella sp. (Fig. 11E)

Remarks: Cephalis large, hemispherical, with a conspicuous apical horn. Thorax thin-walled, conical-inflated, in this specimen the thorax is smaller than L. dictyoceras.

Genus Lithopera ^{Ehrenberg, 1846}

Lithopera neotera Sanfilippo and Riedel (Figs. 11F, 11G and 11H)

Lithopera neotera n.sp. Sanfilippo and ^{Riedel, 1970}, pl. 1, figs. 24-26 - ^{Kamikuri, Motoyama, Nishi & Iwai, 2009}, p. 731, fig. 10AC.

Lithopera renzae Sanfilippo and Riedel (Figs. 11I, 11J, 11K and 11L)

Lithopera renzae n.sp. Sanfilippo and ^{Riedel, 1970}, p. 454, pl. 1, figs. 21-23, 27 - ^{Kamikuri, Motoyama, Nishi & Iwai, 2009}, p. 731, fig. 10T.

Lithopera thornburgi Sanfilippo and Riedel (Fig. 11M)

Lithopera thornburgi n.sp. Sanfilippo and ^{Riedel, 1970}, p. 455, pl. 2, figs. 4-6 - ^{Kamikuri, Motoyama, Nishi & Iwai, 2009}, p. 731, fig. 10 AE.

Genus Lithostrobus ^{Haeckel, 1887}

Lithostrobus hexagonalis Haeckel (Fig. 11N)

Lithostrobus hexagonalis n.sp. ^{Haeckel, 1887}, p. 1475, pl. 79, fig. 20 - ^{Boltovskoy and Riedel, 1987}, p. 100; pl. 5, fig. 11 - ^{Takahashi, 1991}, p. 122, pl. 41, figs. 1-3 - Boltovskoy, 1998, fig. 15.141.

Genus Lophocyrtis ^{Haeckel, 1881}

Lophocyrtis brachythorax (Sanfilippo and

Riedel) (Fig. 11O)

Cyclampterium (?) brachythorax n.sp.

Sanfilippo and ^{Riedel, 1970}, pl. 2, figs. 15-16.

Lophocyrtis (Cyclampterium) brachythorax

Sanfilippo and ^{Riedel, 1970}, p. 457, pl. 2, figs. 1516 - ^{Sanfilippo, 1990}, pl. IV, figs. 4-6 - ^{Kamikuri, Motoyama, Nishi & Iwai, 2009}, p. 731, fig. 13D.

Lophocyrtis tanythorax Sanfilippo and Riedel (Fig. 11P)

Cyclampterium (?) tanythorax n.sp.

Sanfilippo and ^{Riedel, 1970}, pl. 2, figs. 13-14.

Lophocyrtis (Cyclampterium) tanythorax Sanfilippo and ^{Riedel, 1970}, p. 457, pl. 2, figs. 13-14 - Sanfilippo, 1990, pl. 4, figs. 7-10.

Lophocyrtis spp. (Figs. 11Q and 11R)

Remarks: Shell robust, sub-spherical to subellipsoidal in general form. Cephalis sub-spherical, poreless or with a few small pores, with a short apical horn. Thorax inflated, with pores arranged in groups of 8-10, surrounded by a bigger pore. Abdomen thick-walled, with sub-circular pores, it is slightly convex to inflated.

Nassell. gen. and sp. indet. (Figs. 11S and 11T)

Genus Pterocanium ^{Ehrenberg, 1846}

Pterocanium grandiporus (Figs. 12A and 12B) Pterocanium grandiporus n.sp. Nigrini,

1968, p. 57, pl. 1, fig. 7.

Pterocanium praetextum (Haeckel) (Figs. 12C and 12D)

Lychnocanium praetextum n.sp. ^{Ehrenberg, 1872b}, p. 297, pl. 10, fig. 2.

Pterocanium praetextum praetextum (Ehrenberg) - ^{Nigrini and Moore, 1979}, p. N41-N42, pl. 23, fig. 2 - ^{Takahashi, 1991}, p. 115, pl. 36, figs. 15-18.

Pterocanium praetextum group? (Ehrenberg) - ^{Boltovskoy, 1998}, fig. 15.146.

Pterocanium trilobum (Haeckel) (Figs. 12E,

12F, 12G and 12H)

Dictyopodium trilobum n.sp. ^{Haeckel, 1860}, p. 839.

Dictyopodium trilobum^{Haeckel, 1862}, p.

340, pl. 8, figs. 6-10.

Pterocanium trilobum (Haeckel) - ^{Nigrini and Moore, 1979}, p. N45-N46, pl. 23, figs. 4a-c - ^{Boltovskoy, 1998}, p. 78, fig. 15. 145.

Genus Stichocorys ^{Haeckel, 1881}

Stichocorys armata (^{Haeckel) (Figs. 12I and 12J) Cyrtophormis armata n.sp. Haeckel, 1887}, p. 1460, pl. 78, fig. 17.

Stichocorys armata (Haeckel) - Riedel and

Sanfilippo, 1971, p. 1595, pl. 2E, figs. 13-15 - ^{Kamikuri, Motoyama, Nishi & Iwai, 2009}, p. 731, fig. 10O.

Stichocorys delmontensis (Campbell and Clark) (Figs. 12K and 12L)

Eucyrtidium delmontense n.sp. ^{Campbell and Clark, 1944}, p. 56, pl. 7, figs. 19-20.

Stichocorys delmontensis Campbell and

Clark - Sanfilippo and ^{Riedel, 1970}, p. 451, pl. 1, fig. 9 - ^{Kamikuri, Motoyama, Nishi & Iwai, 2009}, p. 731, fig. 10Q.

Stichocorys peregrina (Riedel) (Fig. 12M) Eucyrtidium elongatum peregrinum Riedel, 1953, p. 812, pl. 85, fig. 2.

Stichocorys peregrina^{Riedel - Riedel and Sanfilippo, 1970}, p. 451, pl. 1, fig. 10 - ^{Foreman, 1975}, p. 622, pl. 9, figs. 1-4 - ^{Kamikuri, Motoyama, Nishi & Iwai, 2009}, p. 731, fig. 10P.

Stichocorys sp. (Fig. 12N)

Remarks: Similar to S. peregrina, but the second segment is longer instead to be the third.

Genus Stichopilium ^{Haeckel, 1881}Stichopilium bicorne (Figs. 12O and 12P)

Stichopilium bicorne n.sp. ^{Haeckel, 1887}, p. 1437, pl. 77, fig. 9 - ^{Nigrini and Moore, 1979}, p. N91, pl. 26, figs. 1a-b - ^{Boltovskoy, 1998}, p. 83, fig. 15.161.

Genus Theocorys ^{Haeckel, 1881}

Theocorys (?) spp. (Figs. 12Q, 12R and 12S)

Remarks: Test relatively small, consisting of a cephalis, thorax, and abdomen. Apical robust horn. Pores are distributed through the shell irregularly and dispersed.

Theocorys veneris Haeckel (Fig. 12T) Theocorys veneris Haeckel, 1887 p. 1415, pl. 69, fig. 5 - ^{Boltovskoy and Riedel, 1987}, p. 100, pl. 5, fig. 19 - ^{Takahashi, 1991}, p. 120, pl. 40, figs. 11-14.

Genus Carpocanarium ^{Haeckel, 1887}, emend.

^{Nigrini and Moore, 1979}

Carpocanarium papillosum (Ehrenberg) (Figs. 13A and 13B)

Eucyrtidium papillosum^{Ehrenberg, 1872a},

p. 310-311.

Carpocanarium papillosum (Ehrenberg) - ^{Takahashi, 1991}, p. 132, pl. 45, figs. 16-17.

Dictyocephalus papillosus (Ehrenberg) - ^{van de Paverd, 1995}, p. 255, pl. 76, fig. 24 - ^{Boltovskoy, 1998}, figs. 15.135 a-c.

Carpocanarium aff. papillosum (Figs. 13C and 13D)

aff. Eucyrtidium papillosum Ehrenberg,

1872a, p. 310-311

Remarks: Similar to this species, but rearrangement of the pores and peristome is different.

Family CARPOCANIIDAE^{Haeckel, 1881}, emend. ^{Riedel, 1971a}Genus Carpocanistrum Haeckel, 1887

Carpocanistrum acutidentatum Takahashi (Figs. 13E and 13F)

Carpocanistrum acutidentatum n.sp.

^{Takahashi, 1991}, p. 132, pl. 45, figs. 9, 13-15.

Carpocanistrum spp. (Figs. 13G, 13H and 13I) Remarks: Shell outline sub-spherical to oval. Cephalis indistinguishable from thorax, included within its upper part. Peristome poreless, smooth or provided with teeth. Height of shell: 80-130 μm. (^{Boltovskoy, 1998})

Genus Carpocanopsis ^{Riedel and Sanfilippo, 1971}Carpocanopsis cristata (Carnevale) (Figs. 13J and 13K)

Sethocorys cristata n.sp. ^{Carnevale, 1908}, p. 31, pl. 4, fig. 18.

Carpocanopsis cristatum (Carnevale) - ^{Riedel and Sanfilippo, 1971}, p. 1597, pl. 1G, fig. 16, pl. 2G, figs. 1-7.

Carpocanopsis cristata (Carnevale) - ^{Sanfilippo and Riedel, 1973}, p. 531.

Family CANNOBOTRYIDAE^{Haeckel, 1881}

Genus Botryocyrtis ^{Ehrenberg, 1860a}, b

Botryocyrtis scutum (Harting) (Figs. 13L, 13M and 13N)

Haliomma scutum n.sp. ^{Harting, 1863}, p. 11, pl. 1, fig. 18 - ^{Molina-Cruz, 1977}, p. 338, pl. 6, fig. 14 - ^{Takahashi, 1991}, p. 135, pl. 46, figs. 6-7 - ^{Boltovskoy, 1998}, p. 88, fig. 15. 172.

Botryocyrtis sp. A (Fig. 13O)

Remarks: Eucephalic chamber is enclosed in a rounded cephalis shape. Cylindrical thorax with very small pores. Shell height: ~50 µm. Present two large feelers.

Genus Botryopyle ^{Haeckel, 1881}, emend.^{Petrushevskaya, 1965}

Botryopyle dictyocephalus Haeckel (Figs. 13P and 13Q)

Botryopyle dictyocephalus^{Haeckel, 1887}, p. 1113, pl. 96, fig. 6.

Botryopyle spp. (Figs. 13R and 13S)

Remarks: Large cephalis, from ovoid to pointed shape, thin-walled. Thorax cylindrical, with pores or poreless tube.

Genus Centrobotrys ^{Petrushevskaya, 1965}

Centrobotrys thermophila Petrushevskaya

(Fig. 13T)

Centrobotrys thermophila^{Petrushevskaya, 1965}, p. 115, fig. 20 - ^{Nigrini, 1967}, p. 49, fig. 26, pl. 5, fig. 7 - ^{Boltovskoy, 1998}, p. 88, fig. 15. 175.

Family ARTOSTROBIIDAE^{Riedel, 1967}a in ^{Banner et al., 1967} and Riedel, 1967b, emend. ^{Foreman, 1973}

Genus Artostrobus ^{Haeckel, 1887}

Artostrobus annulatus (Bailey) (Fig. 14A) Cornutella (?) annulata Bailey, 1856, p. 3, pl. 1, figs. 5a-b.

Artostrobus annulatus (Bailey) - ^{Molina-Cruz, 1991}, fig. 2(12-13) - ^{Boltovskoy, 1998}, fig. 15.125.

Genus Botryostrobus ^{Haeckel, 1887}Botryostrobus aquilonaris (Bailey) (Figs. 14B and 14C)

Eucyrtidium aquilonaris n.sp. ^{Bailey, 1856},

p. 4, pl. 1, fig. 9.

Botryostrobus aquilonaris (Bailey) - ^{Nigrini, 1977}, p. 246, pl. 1, fig. 1 - ^{Takahashi, 1991}, p. 128, pl. 44, figs. 9-13 - ^{Boltovskoy, 1998}, p. 84, fig. 15.163.

Botryostrobus auritus (Ehrenberg) (Figs.

14D and 14E)

Lithocampe aurita n.sp. ^{Ehrenberg, 1844}, p. 84.

Artostrobium auritum group (Ehrenberg) - ^{Riedel and Sanfilippo, 1971}, p. 1599, pl. 1H, figs. 5-8.

Lithostrobus lithobotrys (Ehrenberg) - ^{Molina-Cruz, 1977}, p. 336, pl. 7, fig. 17.

Botryostrobus auritus/australis - ^{Boltovskoy, 1998}, p. 84, fig. 15.168.

Botryostrobus bramlettei (Campbell and Clark) (Fig. 14F)

Lithomitra (Lithomitrissa) bramlettei n.sp. ^{Campbell and Clark, 1944}, p. 53, pl. 7, figs. 10-14.

Botryostrobus bramlettei (Campbell and Clark) - ^{Nigrini, 1977}, p. 248, pl. 1, figs. 7-8 - ^{Kamikuri, Motoyama, Nishi & Iwai, 2009}, p. 730, fig. 10G.

Remarks: difference from B. aquilonaris by having a conspicuously swollen third joint, one of important distinguishing characters of species; cephalis small, rather smooth, caplike, not much pitted, apically rounded, and laterally convex.

Genus Phormostichoartus ^{Campbell, 1951}, emend.^{Nigrini, 1977}

Phormostichoartus corbula (Harting) (Figs. 14G and 14H)

Lithocampe corbula n.sp. ^{Harting, 1863}, p. 12, pl. 1, fig. 21.

Phormostichoartus corbula (Harting) - ^{Takahashi and Honjo, 1981}, p. 154, pl. 10, figs. 13-14 - ^{Boltovskoy and Riedel, 1987}, p. 101, pl. 6, fig. 7 - Takahashi, 1991, p. 129, pl. 44, figs. 1416 - Boltovskoy, 1998, p. 84, fig. 15.163.

Phormostichoartus doliolum (Riedel and Sanfilippo) (Fig. 14I)

Artostrobium doliolum n.sp. ^{Riedel and Sanfilippo, 1971}, p. 1599, pl. 1H, figs. 1-3; pl. 8, figs. 14-15.

Phormostichoartus doliolum (Riedel and

Sanfilippo) - ^{Nigrini, 1977}, pl. 1, fig. 14 - Nigrini and Lombari, 1984, p. N181, pl. 31, fig. 5a-b.

Phormostichoartus marylandicus (Martin) (Fig. 14J)

Lithocampe marylandica n.sp. ^{Martin, 1904},

p. 450, pl. 130, fig. 4.

Artostrobium sp. aff. A. doliolum - ^{Riedel and Sanfilippo, 1971}, pl. 1H, fig. 4; pl. 21, figs. 1-8; pl. 3E, figs. 7-9.

Phormostichoartus marylandicus (Martin) - ^{Nigrini, 1977}, p. 253, pl. 2, figs. 1-3.

Genus Siphocampe ^{Haeckel, 1881}, emend.

^{Nigrini, 1977}

Siphocampe spp. A (Figs. 14K and 14L)

Remarks: Specimens present pore rows in the extended part of the abdomen.

Siphocampe arachnea (Ehrenberg) (Fig. 14M)

Lithocampe lineata n.sp. ^{Ehrenberg, 1838}, p. 130.

Eucyrtidum lineatum arachneum^{Ehrenberg, 1861}, p. 299.

Lithomitra lineata group (Ehrenberg) - ^{Riedel and Sanfilippo, 1971}, p. 1600, pl. 1I, figs. 1, 3-11; pl. 21, figs. 14-16.

Siphocampe arachnea (Ehrenberg) - ^{Nigrini, 1977}, p. 255, pl. 3, figs. 7-8.

Siphocampe lineata (Ehrenberg) (Fig. 14N) Lithocampe lineata n.sp. Ehrenberg, 1838, p. 130.

Siphocampe lineata (Ehrenberg) - ^{Nigrini, 1977}, p. 256, pl. 3, fig. 9-10.

Genus Siphostichoartus ^{Nigrini, 1977}Siphostichoartus corona (Haeckel) (Figs. 14O and 14P)

Cyrtophormis (Acanthocyrtis) corona n.sp. ^{Haeckel, 1887}, p. 1462. pl. 77, fig. 15.

Phormostichoartus corona (Haeckel) -

^{Riedel and Sanfilippo, 1971}, p. 1600, pl. 1I, figs. 13-15; pl. 2J, figs. 1-5.

Siphostichoartus corona (Haeckel) - ^{Nigrini, 1977}, p. 257, pl. 2, figs. 5-6.

Genus Spirocyrtis ^{Haeckel, 1881}, emend.

^{Nigrini, 1977}

Spirocyrtis gyroscalaris Nigrini (Fig. 14Q)

Spirocyrtis gyroscalaris n.sp. ^{Nigrini, 1977}, pl. 2, figs. 10-11.

Spirocyrtis gyroscalaris Nigrini - ^{Kamikuri, Motoyama, Nishi & Iwai, 2009}, p. 730, fig. 10E.

Spirocyrtis scalaris^{Haeckel (Figs. 14R and 14S) Spirocyrtis scalaris n.sp. Haeckel, 1887}, p. 1509, pl. 76, fig. 14 - ^{Nigrini, 1977}, p. 259, pl. 2, figs. 12-13 - ^{Takahashi, 1991}, p. 127, pl. 44, figs. 1-2 - ^{Boltovskoy, 1998}, p. 85, fig. 15.166.

Spirocyrtis subtilis Petrushevskaya (Fig. 14T)

Spirocyrtis subtilis n.sp. ^{Petrushevskaya in Petrushevskaya and Kozlova, 1972}, p. 540, pl. 24, figs. 22-24 - ^{Nigrini, 1977}, p. 260, pl. 3, fig. 3 - ^{Kamikuri, Motoyama, Nishi & Iwai, 2009}, p. 730, fig. 10L.

Family PTEROCORYTHIDAE^{Haeckel, 1881},

emend. ^{Riedel, 1967}a in ^{Banner et al., 1967},

emend. ^{Moore, 1972}

Pterocorida ^{Haeckel, 1881}, p. 435

Pterocoryidae ^{Riedel, 1967}a in ^{Banner et al., 1967}, p. 296

Genus Anthocyrtidium ^{Haeckel, 1881}Anthocyrtidium ehrenbergi (Stöhr) (Figs. 15A and 15B)

Anthocyrtis ehrenbergi n.sp. ^{Stöhr, 1880}, p. 100, pl. 3, fig. 21a-b.

Anthocyrtidium ehrenbergi (Stöhr) - ^{Nigrini and Lombari, 1984}, N147, pl. 27, fig. 1.

Anthocyrtidium ophirense (Ehrenberg) (Figs. 15C and 15D)

Anthocyrtis ophirensis n.sp. ^{Ehrenberg, 1872b}, p. 285, pl. 9, fig. 13.

Anthocyrtidium ophirense (Ehrenberg) - MolinaCruz, 1977, p. 337, pl. 6, fig. 10 - ^{Nigrini and Moore, 1979}, p. N67, pl. 25, fig. 1 - ^{Takahashi, 1991}, p. 126, pl. 43, figs. 1-7 - ^{Boltovskoy, 1998}, fig. 15.152.

Anthocyrtidium spp. (Figs. 15E and 15F)

Remarks: Several variations in these specimens. In general consists of a cephalis elongate with a robust three-bladed apical horn. Thorax campanulate, with circular, regular pores arranged longitudinally, peristome distinct within small teeth or undulate. Abdomen is absent.

Genus Calocycletta ^{Haeckel, 1887}

Calocycletta caepa^{Moore (Figs. 15G and 15H) Calocycletta caepa n.sp. Moore, 1972}, p. 150, pl. 2, figs. 4-7 - ^{Kamikuri, Motoyama, Nishi & Iwai, 2009}, p. 736, fig. 14E.

Calocycletta virginis Haeckel (Fig. 15I)

Calocyclas (Calocycletta) virginis n.sp. ^{Haeckel, 1887}, p. 1381, pl. 74, fig. 4.

Calocycletta virginis Haeckel - ^{Riedel and Sanfilippo, 1978}, p. 66, pl. 3, figs. 13-14.

Genus Lamprocyclas ^{Haeckel, 1887}, emend.

^{Nigrini, 1967}

Lamprocyclas junonis (Haeckel) (^{Fig. 15}J) Theoconus junonis n.sp. Haeckel, 1887, p. 1401, pl. 69, fig. 7.

Lamprocyclas junonis (Haeckel) - ^{Caulet, 1986}, pl. 4, fig. 10.

Lamprocyclas maritalis group Haeckel (Figs. 15K and 15L)

Lamprocyclas maritalis n.sp. ^{Haeckel, 1887},

p. 1390, pl. 74, figs. 13-14.

Lamprocyclas maritalis maritalis Haeckel - ^{Nigrini, 1967}, p. 74-76, pl. 7, fig. 5 - ^{Molina-Cruz, 1977}, p. 337, pl. 7, figs. 8-9 - Nigrini and Moore, 1979, p. N75, pl. 25, fig. 4 - ^{Boltovskoy, 1998}, fig. 15.158.

Lamprocyclas sp. A (Fig. 15M)

Remarks: Cephalis sub-spherical, elongate, with a large three-bladed apical horn. Thorax campanulate, thick-walled, with sub-regular, circular pores. Abdomen, with large, sub-regular, circular pores and with sub-terminal teeth.

Genus Lamprocyrtis^{Kling, 1973}

Lamprocyrtis nigriniae (Caulet) (Fig. 15N)

Conarachnium nigriniae n.sp. ^{Caulet, 1971},

p. 3, pl. 3, figs. 1-4; pl. 4, figs. 1-4.

Lamprocyrtis nigriniae (Caulet) - ^{Nigrini and Moore, 1979}, p. N81-N82, pl. 25, fig. 7 - ^{Takahashi, 1991}, p. 127, pl. 43, figs. 17-19 - ^{Boltovskoy, 1998}, p. 82, fig. 15.157.

Lamprocyrtis spp. (Figs. 15O and 15P)

Remarks: Cephalis sub-spherical, with a large three bladed horn. Thorax campanulate, thinwalled, with large polygonal pores increasing in size distally. No abdomen.

Genus Pterocorys ^{Haeckel, 1881}, emend.^{Petrushevskaya, 1971}

Pterocorys minythorax (Nigrini) (^{Fig. 15}Q) Theoconus minythorax n.sp. Nigrini, 1968, p. 57, pl. 1, fig. 8 - ^{Molina-Cruz, 1977}, p. 338, pl. 7, fig. 1.

Pterocorys minythorax (^{Nigrini) - Nigrini and Moore, 1979}, p. N87, pl. 25, fig. 10 - ^{Boltovskoy, 1998}, p. 82, fig. 15.156.

Pterocorys zancleus (^{Müller) (Figs. 15R and 15S) Eucyrtidium zancleum n.sp. Müller, 1855}, p. 672.

Pterocorys zancleus (Müller) - ^{Boltovskoy, 1998}, p. 82, fig. 15.159 - ^{Kamikuri, Motoyama, Nishi & Iwai, 2009}, p. 732, fig. 11R.

Pterocorys sp. (Fig. 15T)

Remarks: Not well preserved specimen.